Comparative development of the crocodylian interclavicle and avian furcula, with comments on the homology of dermal elements in the pectoral apparatus

Abstract

The pectoral apparatus (shoulder girdle plus sternum) of amniotes plesiomorphically includes an unpaired element of dermal origin. In crocodylians, lepidosaurs, and nontherian synapsids (monotremes and their ancestors) this element is identified as the interclavicle, in Testudines (turtles and tortoises) as the entoplastron, and in Aves as the furcula. We investigated embryonic development of the interclavicle in Alligator mississippiensis (American alligator) and of the furcula in Gallus gallus (domestic chicken). The interclavicle and furcula are among the first skeletal elements to ossify, beginning at Ferguson stage 19 (Alligator) and Hamburger and Hamilton stage 33 (Gallus). Both elements: occupy a similar mid-ventral position within the pectoral apparatus; develop from paired (bilateral) cell condensations; never coexist at anytime during ontogeny or in the adult; and undergo intramembranous (i.e., direct) ossification. For both the interclavicle and the furcula, the initial onset of ossification is concomitant with mineralization of elements of the dermatocranium, and occurs in advance of mineralization of the replacement bones (e.g., scapula, metacoracoid) of the pectoral apparatus. Shortly after the initiation of ossification the paired condensations of both elements fuse. For each of Alligator and Gallus, only one pair of skeletogenic condensations is present during embryonic development. Based on these data and a review of the evolution and development of dermal elements in the pectoral apparatus, we conclude that the interclavicle is equally parsimonious as a homolog of the furcula.